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      Phenotypic and developmental plasticity of xylem in hybrid poplar saplings subjected to experimental drought, nitrogen fertilization, and shading

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          Abstract

          Variation in xylem structure and function has been extensively studied across different species with a wide taxonomic, geographical, and ecological coverage. In contrast, our understanding of how xylem of a single species can adjust to different growing condition remains limited. Here phenotypic and developmental plasticity in xylem traits of hybrid poplar ( Populus trichocarpa× deltoides) was studied. Clonally propagated saplings were grown under experimental drought, nitrogen fertilization, and shade for >30 d. Xylem hydraulic and anatomical traits were subsequently examined in stem segments taken from two different vertical positions along the plant’s main axis. The experimental treatments affected growth and development and induced changes in xylem phenotype. Across all treatments, the amount of leaf area supported by stem segments (A L) scaled linearly with stem native hydraulic conductivity ( K native), suggesting that the area of assimilating leaves is constrained by the xylem transport capacity. In turn, K native was mainly driven by the size of xylem cross-sectional area (A X). Moreover, the structural and functional properties of xylem varied significantly. Vulnerability to cavitation, measured as the xylem pressure inducing 50% loss of conductivity (P50), ranged from –1.71MPa to –0.15MPa in saplings subjected to drought and nitrogen fertilization, respectively. Across all treatments and stem segment positions, P50 was tightly correlated with wood density. In contrast, no relationship between P50 and xylem-specific conductivity ( K S) was observed. The results of this study enhance our knowledge of plant hydraulic acclimation and provide insights into common trade-offs that exist in xylem structure and function.

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          Towards a worldwide wood economics spectrum.

          Wood performs several essential functions in plants, including mechanically supporting aboveground tissue, storing water and other resources, and transporting sap. Woody tissues are likely to face physiological, structural and defensive trade-offs. How a plant optimizes among these competing functions can have major ecological implications, which have been under-appreciated by ecologists compared to the focus they have given to leaf function. To draw together our current understanding of wood function, we identify and collate data on the major wood functional traits, including the largest wood density database to date (8412 taxa), mechanical strength measures and anatomical features, as well as clade-specific features such as secondary chemistry. We then show how wood traits are related to one another, highlighting functional trade-offs, and to ecological and demographic plant features (growth form, growth rate, latitude, ecological setting). We suggest that, similar to the manifold that tree species leaf traits cluster around the 'leaf economics spectrum', a similar 'wood economics spectrum' may be defined. We then discuss the biogeography, evolution and biogeochemistry of the spectrum, and conclude by pointing out the major gaps in our current knowledge of wood functional traits.
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            Size and function in conifer tracheids and angiosperm vessels.

            The wide size range of conifer tracheids and angiosperm vessels has important consequences for function. In both conduit types, bigger is better for conducting efficiency. The gain in efficiency with size is maximized by the control of conduit shape, which balances end-wall and lumen resistances. Although vessels are an order of magnitude longer than tracheids of the same diameter, they are not necessarily more efficient because they lack the low end-wall resistance of tracheids with torus-margo pits. Instead, vessels gain conducting efficiency over tracheids by achieving wider maximum diameters. End-walls contributed 56-64% to total xylem resistance in both conduit types, indicating that length limits conducting efficiency. Tracheid dimensions may be more limited by unicellularity and the need to supply strength to homoxylous wood than by the need to protect against cavitation. In contrast, the greater size of the multicellular vessel is facilitated by fibers that strengthen heteroxylous wood. Vessel dimensions may be most limited by the need to restrict intervessel pitting and cavitation by air-seeding. Stressful habitats that promote narrow vessels should favor coexistence of conifers and angiosperms. The evolution of vessels in angiosperm wood may have required early angiosperms to survive a phase of mechanic and hydraulic instability.
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              Scaling of angiosperm xylem structure with safety and efficiency.

              We tested the hypothesis that greater cavitation resistance correlates with less total inter-vessel pit area per vessel (the pit area hypothesis) and evaluated a trade-off between cavitation safety and transport efficiency. Fourteen species of diverse growth form (vine, ring- and diffuse-porous tree, shrub) and family affinity were added to published data predominately from the Rosaceae (29 species total). Two types of vulnerability-to-cavitation curves were found. Ring-porous trees and vines showed an abrupt drop in hydraulic conductivity with increasing negative pressure, whereas hydraulic conductivity in diffuse-porous species generally decreased gradually. The ring-porous type curve was not an artifact of the centrifuge method because it was obtained also with the air-injection technique. A safety versus efficiency trade-off was evident when curves were compared across species: for a given pressure, there was a limited range of optimal vulnerability curves. The pit area hypothesis was supported by a strong relationship (r2 = 0.77) between increasing cavitation resistance and diminishing pit membrane area per vessel (A(P)). Small A(P) was associated with small vessel surface area and hence narrow vessel diameter (D) and short vessel length (L)--consistent with an increase in vessel flow resistance with cavitation resistance. This trade-off was amplified at the tissue level by an increase in xylem/vessel area ratio with cavitation resistance. Ring-porous species were more efficient than diffuse-porous species on a vessel basis but not on a xylem basis owing to higher xylem/vessel area ratios in ring-porous anatomy. Across four orders of magnitude, lumen and end-wall resistivities maintained a relatively tight proportionality with a near-optimal mean of 56% of the total vessel resistivity residing in the end-wall. This was consistent with an underlying scaling of L to D(3/2) across species. Pit flow resistance did not increase with cavitation safety, suggesting that cavitation pressure was not related to mean pit membrane porosity.
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                Author and article information

                Journal
                J Exp Bot
                J. Exp. Bot
                jexbot
                jexbot
                Journal of Experimental Botany
                Oxford University Press (UK )
                0022-0957
                1460-2431
                November 2012
                23 October 2012
                23 October 2012
                : 63
                : 18
                : 6481-6491
                Affiliations
                University of Alberta , Department of Renewable Resources , 4–42 Earth Sciences Building, Edmonton, AB, Canada, T6G 2E3
                Author notes
                * To whom correspondence should be addressed. E-mail: lenka.plavcova@ 123456ualberta.ca
                Article
                10.1093/jxb/ers303
                3504499
                23095999
                a40b2914-53a8-4c07-a686-e34784ed587c
                © 2012 The Author(s).

                This is an Open Access article distributed under the terms of the Creative Commons Attribution Non-Commercial License ( http://creativecommons.org/licenses/by-nc/2.0/uk/) which permits unrestricted non-commercial use, distribution, and reproduction in any medium, provided the original work is properly cited.

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                Page count
                Pages: 11
                Categories
                Research Paper

                Plant science & Botany
                cavitation,wood density,vessels,xylem embolism.,hydraulic conductivity,phenotypic plasticity

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